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APPLIED ODONATOLOGY

A review of odonatology in freshwater applied ecology and conservation science

Jason T. Bried1,3 and Michael J. Samways2,4

1Department of Integrative Biology, Oklahoma State University, Stillwater, Oklahoma 74078 USA 2Department of Conservation Ecology and Entomology, Stellenbosch University, Matieland 7602 South Africa

Abstract: The academic study of dragonflies and damselflies (odonatology) is well established, but relatively limited attention has been given to odonates in the context of applied ecology and conservation science. We used the Web of Science™ and Odonatological Abstract Service (ISSN 1438-0269) to capture trends in primary literature, characterize study features (habitats, life stages, etc.), identify research themes, and suggest future directions for odonatology in freshwater applied ecology and conservation science. We found no papers in this area prior to 1980, and 411 papers from 1980 through 2013. Nearly 75% of these papers were recent (since 2005) and >40% were very recent (since 2010). We identified several broad and overlapping research themes: 1) model taxa, 2) tools and indicators, 3) odonate-centered work, and 4) methodological issues and improvements (field sampling, data modeling/simulation, conservation/landscape-scale genetics). We found more reliance on field- based observational approaches than experiments and model-driven exercises, although the number of papers using model-driven exercises is rapidly increasing. We found a strong focus on adult stages, odonate assem- blages, the as a whole, and studies of particular species. We identified research priorities in areas such as ecological valuation and management, monitoring and assessment, climate change and landscape planning, concordance with other taxa, effects of urbanization, data modeling/simulation, and rare-species ecology and conservation. To help establish an identity and facilitate communication, we suggest naming this diverse realm “applied odonatology”. We think applied odonatology has a good future for a range of topics from conservation genetics and population ecology to assessments of anthropogenic impacts and the conservation of biodiversity. Key words: dragonflies, assessment, climate change, monitoring, landscape planning, freshwater health, biodi- versity conservation

The academic study of dragonflies and damselflies (odo- water applied ecology and conservation science. No gen- natology) is well established. However, relatively few odo- eral review of odonatology has been done in this broad natologists work outside of and systematics, context, although detailed reviews have been made of behavioral ecology, evolutionary ecology, and other prom- relevant subject areas (e.g., Hassall and Thompson 2008). inent areas of classical research. The major books (Corbet Our purpose was to review the odonatological literature 1999, Córdoba-Aguilar 2008), treatises (e.g., Corbet 1980, for studies related to applied ecology and conservation Stoks and Córdoba-Aguilar 2012), and flagship journals of science, and to provide a formal synthesis and baseline odonatology reveal a productive legacy, but also relatively for assessing advances in freshwater applied science and limited attention to matters of applied ecology and con- conservation via odonates. Rather than diving deeply into servation science. For example, <5% of the 275 papers any particular subject area, we tried to cover many dif- published during 2009–2013 in Odonatologica and the In- ferent areas and provide a general map of the diverse ternational Journal of Odonatology are clearly relevant to field. We excluded nonscientific topics, such as odonates applied ecology or conservation science. Nevertheless, the in ecotourism, culture, and symbolism, and environmen- relatively few odonatologists working in this area and tal education, despite the importance of these areas to researchers based in other fields who often or sometimes public awareness and conservation (Primack et al. 2000, use odonates as principal study subjects are making im- Lemelin 2007). We also did not consider the extensive portant contributions. work in odonate faunistics as being inherently applied, A growing body of dragonfly-related research deals even though faunistic efforts provide essential informa- with the study or use of odonates in the context of fresh- tion for assessments of conservation status and niche mod-

E-mail addresses: [email protected]; [email protected]

DOI: 10.1086/682174. Received 10 April 2014; Accepted 25 November 2014; Published online 19 May 2015. Freshwater Science. 2015. 34(3):1023–1031. © 2015 by The Society for Freshwater Science. 1023 1024 | Applied odonatology J. T. Bried and M. J. Samways eling. Our goal was to reach out to odonatologists and the Gee et al. 1997, Maxted et al. 2000, King and Richardson broader freshwater-science community seeking tools and 2007). However, we did count studies of broader issues model taxa with which to address pressing issues in the that used odonates and other taxa as model or target realm of applied ecology and conservation science (Strayer groups (e.g., Siegfried 1993, Richter et al. 1997, Palmer 2006). 1999, Le Viol et al. 2009, Rosset et al. 2013, 2014). We used the Odonatological Abstract Service (ISSN 1438-0269) to expand the Web of Science search results. METHODS The Odonatological Abstract Service, published by the In- We extracted odonatology literature stored in the Web ternational Dragonfly Fund in cooperation with the World- of Science™ by querying TOPIC: (dragonfl*ORdamselfl* wide Dragonfly Association, is an ongoing effort to com- OR Anisoptera OR Zygoptera OR Odonata) AND YEAR: pile odonate-related publications (in any language) using (1900–1980 at 5-y intervals, 1980–2013 annually). We Google searches, publisher databases, alerts from Google searched for primary literature (international peer-reviewed and publishers,® and a correspondence network of odo- journals) that appeared to meet the criteria for applied eco- natologists. This repository contains the world’slargestcol- logical or conservation science, as defined by the aims, lection of odonate-related literature, including items from scope, and content of several leading journals (e.g., Aquatic primary and regional/domestic journals, museum bulletins, Conservation: Marine and Freshwater Ecosystems, Biolog- government reports and technical series, conference ab- ical Conservation, Conservation and Diversity, Jour- stracts and proceedings, theses and dissertations, and other nal of Applied Ecology). We assessed each candidate paper sources. We viewed titles, keywords, abstracts, and com- and categorized it relative to the 17 subject areas listed in piler annotations of all primary literature available from Table 1. The small number of applied/conservation papers 1997 (when the abstract service began) through 2013, and that did not clearly fit at least one of the main subject stopped at the June 2014 issue. We avoided papers in headings were classified as Miscellaneous. We excluded recently launched open-access journals because of con- purely faunistic or nonscientific surveillance investigations cerns about credibility (Bohannon 2013). The complete ser- and the many studies in which results were reported for vice was accessed using a fully searchable database built and odonates but that did not explicitly target odonates (e.g., maintained by M. Schorr (International Dragonfly Fund).

Table 1. Major odonatology research areas in freshwater applied ecology and conservation science using the Web of Science™ (1980–2013) and Odonatological Abstract Service (1997–2013), with the total number of articles from international peer-reviewed journals (No. papers) and frequently associated topics. Categories are not mutually exclusive; many entries were classified under 2 or 3 subject areas (see Appendix S1). Subject area No. papers Key research topics

Biocontrol 16 Mosquitos, rice pests Conservation status 23 Priority species, status ranks, vulnerability, Red Lists Disturbance and threats Climate change 38 Range shifts, phenologic shifts Invasives 19 Introduced fish, zebra mussels, Acacia trees Urbanization 13 City landscapes, gradients of human development, secondary habitats Other 65 Agroecology, forestry, altered vegetation Diversity and distributions 36 Biodiversity, biogeography, hotspots Ecological management 34 Habitat creation, recovery, remediation, restoration Genetics 36 Conservation genetics, landscape-scale genetics Landscape ecology 39 Movement and dispersal, metapopulations and metacommunities, fragmentation Methods and modeling 49 Field surveys, genetic methods, climate and niche modeling Monitoring and assessment Indicators 57 Pollution, climate change, diversity, ecological condition, fluctuating asymmetry Other 44 Bioassessment, index applications, population and community trends Planning and valuation 47 Reserve selection/design, species recovery plans, secondary habitats Pollution and toxicology 60 Agriculture/pesticides, heavy metals, developmental instability Rare species ecology 26 Autecology, habitat requirements, population trends Miscellaneous 6 Various topics Volume 34 September 2015 | 1025

For each paper we recorded the year, journal name, geographic location, study type, major habitat, life stage, community vs species focus, general research theme (de- scribed below in ‘General research themes’), and up to 3 subject areas (see Table 1) based on perceived rele- vance. The categories in several information fields were not mutually exclusive, and many papers fit into >1 cate- gory. We left fields blank when the information was lack- ing or ambiguous in the abstract or not logically or sci- entifically applicable—e.g., the habitat type in laboratory studies and in some modeling exercises. Geographic loca- tion was the study location (when applicable and discern- ible) or the location of the corresponding author’s institu- tion. We viewed the full-text articles as needed, using an extensive library of >13,200 portable document files com- piled by M. Schorr, journals in library holdings, and inter- library loan service at Oklahoma State University. Figure 1. Cumulative odonate literature (papers in interna- All database searches were conducted by one of us tional peer-reviewed journals, excluding recently launched (JTB), and all the numbers and figures reported in our open-access journals) in the realm of applied ecology and con- servation science based on the Web of Science (1980–2013) paper are best interpreted as minimum estimates. The 2nd ™ and complete Odonatological Abstract Service (1997–2013). author (MJS) checked the various classifications for 30% of fi We found no odonate papers on applied ecology or conserva- the selected papers and did not nd any disagreements. tion science before 1980. We focused on primary literature because the full text is more readily available than for gray literature and to cap- ture a globally representative cross-section of the broadest ture prominently in some areas (e.g., aquatic toxicology), trends shaping applications and conservation in odonatol- but contributions were lacking in newer applied research ogy. We acknowledge that regional variation exists in re- areas, such as climate change, invasive species, conservation search priorities, and we recognize that a vast amount of genetics, urbanization, and restoration ecology. Conservation- information on odonates is contained in domestic/regional status assessments, including regional and global Red Lists, journals and other gray literature (M. Schorr, personal became more frequent during the 1980s and 1990s, as did communication). evaluations of odonates for use in biocontrol of mosquitos (e.g., Miura et al. 1990, Sebastian et al. 1990). Total contri- butions increased steadily into the 2000s (Fig. 1), but the RESULTS percentage of papers on applied ecology and conservation We found no applied ecological or conservation sci- science remained low (<5%). ence entries (searching primary literature only) in the Web Empirical, observation-based research has been more of Science and Odonatological Abstract Service prior to common than experiments (Fig. 2A), suggesting a preva- 1980 (searching at 5-y intervals starting from 1900), and lence of fieldwork, although experimental manipulations 411 papers in the annual search from 1980 through 2013 and laboratory trials occur regularly in odonate-based bio- (Appendix S1). International peer-reviewed journal arti- control and ecotoxicology (e.g., Hardersen et al. 1999, Singh cles constituted ∼35% of the total English-language entries et al. 2003). The growing number of modeling and simula- (∼8800) in the Odonatological Abstract Service. We treated tion efforts, although a small fraction overall (Fig. 2A), is a several journal special series with an applied/conservation significant addition to the experiments and observational theme as single records. The number of new papers per approaches in odonatology. Attention to conservation ge- year generally increased during the 34-y period (Fig. 1). netics and landscape-scale genetics has increased greatly Nearly 75% of papers appeared after 2005 and >40% after over the past decade. 2010. Studies were done most frequently at ponds or wet- Research characteristics of the 411 papers are given in lands, least frequently at lakes, and with intermediate fre- Appendix S1. Here we provide an overview. The first clear quency at rivers and streams (Fig. 2B). This pattern was records of applied ecology or conservation science (Moore partly driven by the breeding habitats of target species. 1980, Garrison and Hafernik 1981) focused on population Adult and larval stages were used far more frequently than ecology, habitat requirements, and threat factors of 2 rare eggs and exuviae (Fig. 2C), which were surveyed more often damselfly species, and were followed by 6 more papers dur- in combination with other stages (6 and 23 papers, respec- ing the 1980s (Fig. 1). At this time, odonates began to fea- tively) than alone (2 and 6 papers). Last, a community-level 1026 | Applied odonatology J. T. Bried and M. J. Samways

Figure 2. Distributions of study types (A), major habitats (B), life stages (C), and community vs species focus (D) in odonatology application and conservation science based on entries in the Web of Science™ (1980–2013) and complete Odonatological Abstract Service (1997–2013). In panel A, most ‘other’ papers were focused on assessments of conservation status. In panel B, ‘other’ papers included miscellaneous study habitats, such as springs, ditches, uplands, and phytotelmata. About 10% of the wetland studies were done in rice paddies, and ∼5% of the pond studies involved artificial mesocosms. focus was at least as common as target-species research, with Odonates are excellent models with which to test ideas, 215 vs 186 papers, respectively (Fig. 2D). problems, and theory in applied landscape and commu- nity ecology. For example, they have been used to address GENERAL RESEARCH THEMES questions about genetics, life history, and movement dy- We propose several research themes broadly defined as: namics in human-dominated heterogeneous landscapes 1) odonates as model organisms, 2) tools and indicators, (9, 18, 48, 51, 56, 104, 105, Feindt et al. 2014, Harms et al. 3) odonate-centered work, and 4) methodology (as a pri- 2014, Suhonen et al. 2014). Other applications include the mary or secondary focus) (Table 2). Odonate-centered work study of metapopulation and metacommunity dynamics in comprised the largest number of database entries (173 pri- patchy or changing environments (16, 82, 99), partitioning mary publications) followed by tools and indicators (154), variation in community responses among natural and an- model organisms (104), and methods (59, ∼50 with pri- thropogenic drivers (65, 82), and testing predictions or mary focus on methods or modeling). The categories are implications of island biogeographic theory for biodiver- not mutually exclusive, and at least 18.5% of the 411 pa- sity conservation (17, 83, 86, 121, Heiser et al. 2014). pers fit in multiple themes. In the discussion below, we This category also includes studies of odonate behav- cite articles published before 2014 as numerals and list ior in the context of major environmental stressors, such them in Appendix S2. as predicting how pesticides, invasive species, and cli- mate change may alter growth or predator–prey interac- tions (47, 96, 102, 112, 119). Many experimental studies Odonates as model taxa of tadpole antipredator defense were done with odonate Authors of papers in this category used odonates as larvae as the model predator, and some investigators ex- principal subjects in studies designed to address broader plicitly incorporated applied perspectives, such as pollu- problems in applied ecology and conservation science. tion stress and invasion biology (55, 103, 108, 109, 118). Volume 34 September 2015 | 1027

Table 2. General research themes for odonatology in freshwater applied ecology and conservation science. Subject areas correspond to Table 1. Values in parentheses are the percentage of papers in that subject area across the 4 categories (using single-category papers only). Category Main subject areas

1. Model taxa for applied ecology and conservation science Climate change (64%), landscape ecology (53%) 2. Tools and indicators for conserving other taxa and systems Indicators (93%), biocontrol (93%) 3. Odonate-centered, concern for species and their habitats Rare species ecology (91%), conservation status (86%) 4. Method issues and improvement Methods and modeling (93%)

More broadly, authors of these studies used odonates to 6, 12, 64, Hart et al. 2014). Odonates also can be used when help test whether predators modify the effects of nutrients/ evaluating biodiversity in secondary or man-made aquatic contaminants or invasive species on aquatic communities habitats (e.g., 61, 99) or may have potential as a cross- (116, 123). Odonates have good potential as model systems taxon surrogate to represent broader freshwater diversity for aquatic toxicology (Stoks et al. 2015), including the trans- patterns (13, 19, 25, 32, 38, 94, 122). Many other poten- fer of toxic materials to higher taxa or across the land–water tial uses of odonates as indicators, such as in conserva- boundary (1, 106). As mid-level consumers with complex tion genetics applied to environmental monitoring (26) or life histories, odonates may have a key role in trophic cas- when delineating protective buffer zones adjacent to hab- cades addressed from the perspective of invasive species itat patches (29), have received little attention. (92), local habitat alteration (62), and large-scale environ- Scientific consideration of odonates for biocontrol con- mental change (98). tinues to be common and probably is the best example of Other types of research done with odonates as model using odonates as agents of ecological services to humans taxa include studies of biologically based selection and (54). Much of the research on using odonates for biocon- design of reserves (81, 85); shifts in phenology, distribu- trol takes place in India and is focused on mosquitos (21, tion, and diversity in response to modern climate change 35, 110), but odonate-based pest control has been evalu- (27, 36, 45, 115, Li et al. 2014); and planning of reserve net- ated in other contexts (3, 15, 46). Authors of recent reviews works under future climate scenarios (Bush et al. 2014a). in biocontrol have highlighted the predatory capacity of The line between model taxa and tools/indicators (next sec- odonates and called for further research (37, 63). Another tion) may blur in these studies. example of service provided by odonates is conservation work driven by nonscientists but built on scientificunder- Odonates as tools and indicators pinnings (e.g., 10, 14). Studies that most clearly belong in this category are those that were done to develop or test odonate-based Odonate-centered research indices for freshwater monitoring and evaluation, such as Research in this category generally does not reach the Dragonfly Biotic Index (38, 87) and a habitat associa- broader conclusions and implications for other taxa (in- tion index for river–floodplain systems (11, 20, Chovanec cluding humans) or for ecosystem health and functioning. et al. 2014). More broadly, odonates often are viewed as Instead, the studies are intrinsically motivated and tar- indicators or sentinels of environmental change and per- geted at particular species or assemblages (e.g., 60, 70), turbation to freshwaters (52, 74, 80, 117). For example, covering topics, such as rankings of conservation status, indicator studies may use odonate mortality levels and threat assessments, distributions and vulnerability, levels developmental instability to assess pesticides and habitat of endemism, and conservation genetics. Recent accom- alteration (e.g., 2, 5, 34, 107) and the potential transfer of plishments include broad status assessments and prioritiza- contaminants to higher taxa and terrestrial systems (e.g., tion schemes across regions, nations, continents, and the 1, 40, 49). Most evidence supporting odonates as indi- globe (e.g., 31, 33, 57, 67, 69, 93, 124, White et al. 2015). At cators currently is context-specific and related to partic- the same time, finer-grained status rankings (e.g., 120) rec- ular stressors or water-quality variables rather than to gen- ognize the need to account for local rarity patterns and the eral ecological condition of freshwaters and surrounding logistical/jurisdictional realities of implementing conserva- landscapes (Kutcher and Bried 2014). tion actions. Threat assessments may range from particular Odonates may be used as indicators for monitoring aspects of odonate biology (e.g., 89) to use of broad ecologi- habitat creation, improvement, and restoration (8, 22, 50, cal patterns to guide conservation and management plans 59) or as a focal group in prioritization schemes, iden- (e.g., 39). Odonatologists have studied odonate responses tification of biodiversity hotspots, and assigning value to to a variety of anthropogenic threats, such as forestry and natural areas for inclusion in parks or protected areas (e.g., agriculture (e.g., 7, Koch et al. 2014), mining (e.g., 78), 1028 | Applied odonatology J. T. Bried and M. J. Samways aquatic invasions (e.g., 4, 58), boating (e.g., Hall et al. 2015), odonatology (90). Controlling for imperfect detectability is and urbanization (e.g., Monteiro-Júnior et al. 2014). Many important for estimating occupancy and abundance from odonate-centered projects have focused on the genetics or standardized field surveys and may help investigators ex- ecology of rare and declining species (e.g., 72, 114, Dolný ploit extensive opportunistic data (71, 77, 125). et al. 2014, Monroe and Britten 2014), and certain species of concern, such as Coenagrion mercuriale in Europe, He- miphlebia mirabilis in Australia, hirosei in CONCLUSIONS AND RESEARCH PRIORITIES Japan, and Somatochlora hineana intheUSA,havereceived Applied ecology and conservation science involving odo- considerable attention. nates has undoubtedly been shaped by greater popular and scientific awareness for insect conservation in general (Sam- ways 1994, 2005, 2007, Bassett et al. 2009, New 2012). How- Methodology ever, despite significant progress, the literature of odo- The importance of methodology is strongly appreciated natology in freshwater applied ecology and conservation throughout the biological sciences, as exemplified by nu- science is sparse relative to the literature in behavioral ecol- merous textbooks (e.g., 76) and journals, such as Methods ogy, sexual selection, morphological and molecular sys- in Ecology and Evolution. We found 49 papers focused tematics, and other prominent areas (see Córdoba-Aguilar largely or exclusively on methods and modeling (Table 1). 2008). Our broad circumscription in Table 2 recognizes Authors of such papers view methods as a means to an the intrinsic and utilitarian value of odonates and the im- end and as an end in itself and strive to develop new portance of underlying methods. Research in applied ecol- methods, improve existing ones, or evaluate alternatives. ogy and conservation science tends to be more interdis- Papers in this category often are specifically about meth- ciplinary and integrative than basic research, especially in ods for studying odonates, but broader methods issues studies of model taxa and tools/indicators where odonates can be addressed using odonates (e.g., 79, 95, Yoshioka usually are not the ultimate concern. These themes help et al. 2014). place odonatology into the wider context of freshwater ap- Papers in this category tend to address genetic methods, plied ecology and conservation science. field methods, and data modeling/simulation approaches. The results of our literature review indicate that de- With regard to genetics, applications have ranged from scriptive or comparative observational approaches are developing genetic markers for conservation and popula- used more than natural or manipulative controlled ex- tion studies of vulnerable species (e.g., 24, 88) to new statis- periments and modeling approaches, but the use of mod- tical approaches for quantifying landscape genetic structure eling is increasing rapidly and can help compensate for (113). Other examples include nonlethal tissue sampling for deficiencies in purely observational studies, such as im- protected species (28, 73), comparing genetic vs field es- perfect detectability. Experiments in odonatology have re- timates of population size and dispersal rates (42, 43), lied more heavily on larvae (easier to manipulate than efficacy of DNA extraction from museum-archived mate- adults) and particular species, whereas the observational rial (44), and DNA barcoding to identify population bound- approach seems to favor adult stages and species assem- aries and conservation units (53). blages. Wetlands appear to be popular study habitats, but Field-methods research deals with principles of study we caution that the term wetlands is often misused to refer design (sample size, representativeness, stratification, etc.), to aquatic systems that are not bogs, marshes, swamps, or sampling logistics, statistical efficiency (minimizing vari- other true wetland types (Mitsch and Gosselink 2007). ance), and observation biases. For example, investigators Moreover, investigators may have sampled wetland hab- have evaluated line–transect distance sampling (30) and itats within an encompassing aquatic system (e.g., a lake- mark–recapture estimates of dispersal and population size fringe wetland) and named the habitat type according to (41, 101, Harms et al. 2014). Others have assessed the util- the sampled area and not the larger habitat complex. ity of exuvial sampling for sensitive species (23), the trade- Our review suggests several research areas that would offsofsamplingdifferent life stages (75, 90, 97), and em- benefit from more attention. These are areas where odo- pirical guidelines for cost-effective field surveys (91). nates could make a substantial contribution, both as sub- Researchers are active in building and validating odo- jects in their own right and as tools or models for other nate habitat and niche models, which are being used in- taxa and ecological systems. The general priorities in- creasingly for predicting climate-induced range shifts and clude, but are not limited to: life-history alterations (66, 84, 111, Bush et al. 2014b). As odonate species distribution modeling becomes more main- • Environmental change: synergies between climate stream, critical evaluations of ecological assumptions and change and landscape/habitat alteration, geographi- model accuracy will be needed (68, 100, Collins and Mc- cal range shifts, assisted dispersal strategies (Hoegh- Intyre 2015). Modeling the factors that control variation in Guldberg et al. 2008, Loss et al. 2011); e.g., How probability of species detection deserves greater attention in resilient are odonates to climate change? Volume 34 September 2015 | 1029

• Reserve/natural area planning: protected area buf- Viability Analysis), captive rearing and species rein- fer zones, development of large-scale reserve net- troduction, conservation/landscape-scale genetics; works, spatiotemporal congruence with other taxa; e.g., Can we afford to address key threats to certain e.g., How well do Biosphere Reserves function? Are odonate species (fine-filter conservation) or should odonates surrogates for freshwater biodiversity? we operate only at the community and ecosystem • Ecological valuation and management: cost–benefit levels (coarse-filter)? analyses of management interventions (sensu Mor- lando et al. 2012), ecological services including bio- Odonates have inherent strengths for use in fresh- control of mosquitos and agricultural pests, value water applied ecology and conservation science, including of natural vs created habitats, value of secondary ontogenetic linkage of aquatic and terrestrial systems, sen- habitats; e.g., Are rare or threatened species con- sitivity to thermal conditions and water pollution in gen- served in secondary habitats? Can odonates be used eral, rapid response to large-scale environmental change, more widely for biocontrol of mosquitoes and, if complex trophic interactions, keystone status in some ephem- so, how might the habitat (such as rice paddies) be eral or fishless systems, and charisma to engage popular modified to accommodate them? interest. We propose that dragonflies and damselflies are • Urbanization and development: value of urban ponds valuable as model taxa, tools and indicators, and subjects for conservation of odonates, importance of ripar- in their own right. We suggest referring to the academic ian corridors in urban settings, road ecology; e.g., study or science-based usage of odonates in the realm of Is there a certain number, size distribution, or spa- applied ecology and conservation science as applied odo- tial distribution of ponds and wetlands that will natology, a convenient name representing a diverse yet dis- maximize odonate diversity in urban settings? Are tinctive subset of odonatology. As research accumulates riparian zones important connectivity features in in priority subject areas, applied odonatology will require developed areas? Are odonates as threatened as targeted reviews and meta-analyses to synthesize the im- butterflies by roads? Do we assume that all roads portant trends and facilitate further advancement. We think are barriers/obstacles, or are roads with low traffic applied odonatology has a good future for a range of topics volume flight corridors? from conservation genetics and population ecology through • Monitoring and modeling: community-level trends to assessing anthropogenic impacts and the conservation analysis (could use hierarchical occupancy mod- of biodiversity. els; DeWan and Zipkin 2010), estimating species tolerance values (to pollution or anthropogenic stress in general), consequences of native and nonna- ACKNOWLEDGEMENTS tive species invasions, tracking restoration progress We are grateful to Martin Schorr for sharing his massive Odo- based on habitat potential (Bried et al. 2014, Cho- nata database and personal library and for helping to compile vanec et al. 2014), field validation of distribution some of the statistics. Guest Editor Chris Hassall and 2 anon- model predictions, accounting for imperfect and ymous reviewers provided feedback that led to major improve- varying detection probability, inferences from ex- ment of the original manuscript. MJS acknowledges the Na- fi tensive citizen-science data sets; e.g., Can dragon- tional Research Foundation, South Africa, for nancial support. fly-based ecological integrity indices be applied at large spatial scales (a pan-African Freshwater Health fl LITERATURE CITED Assessment based on the Dragon y Biotic Index Basset, Y., B. A. Hawkins, and S. R. Leather. 2009. Visions for is being done now)? Which odonate metrics (e.g., insect conservation and diversity: spanning the gap between community structure vs ecological traits) work best, practice and theory. Insect Conservation and Diversity 2:1–4. and do odonates add value to the Ephemeroptera, Bohannon, J. 2013. Who’s afraid of peer review? Science 342:60– Plecoptera, Trichoptera trio of sentinels? Can citi- 65. zen scientists be involved in recording particularly Bried, J., T. Tear, R. Shirer, C. Zimmerman, N. Gifford, S. sensitive species? Campbell, and K. O’Brien. 2014. A framework to integrate • Field methods: optimal sampling design (site vs habitat monitoring and restoration with endangered insect re- – survey replication tradeoff ), survey effort guide- covery. Environmental Management 54:1385 1398. lines (frequency, duration, seasonal placement), in- Bush, A., V. Hermoso, S. Linke, D. Nipperess, E. Turak, and L. Hughes. 2014a. Freshwater conservation planning under cli- terchangeability of major life stages (mature adults, mate change: demonstrating proactive approaches for Aus- tenerals, exuviae, larvae), criteria to extract locally tralian Odonata. Journal of Applied Ecology 51:1273–1281. autochthonous (resident) species in adult surveys; Bush, A. A., D. A. Nipperess, D. E. Duursma, G. Theischinger, E. e.g., Can we establish universal procedures? Turak, and L. Hughes. 2014b. Continental-scale assessment • Rare species ecology and conservation: brood size of risk to the Australian Odonata from climate change. PLoS estimation, prediction of extinction risk (Population ONE 9:e88958. 1030 | Applied odonatology J. T. Bried and M. J. Samways

Chovanec, A., M. Schindler, J. Waringer, and R. Wimmer. 2014. Hoegh-Guldberg, O., L. Hughes, S. McIntyre, D. B. Lindenmayer, The Dragonfly Association Index (Insecta: Odonata)—A tool C. Parmesan, H. P. Possingham, and C. D. Thomas. 2008. for the type-specific assessment of lowland rivers. River Re- Assisted colonization and rapid climate change. Science 321: search and Applications (in press). doi:10.1002/rra.2760 345–346. Collins, S., and N. E. McIntyre. 2015. Modeling the distribution King,R.S.,andC.J.Richardson.2007.Subsidy-stressresponseof of odonates: a review. Freshwater Science 34:1144–1158. macroinvertebrate community biomass to a phosphorus gradi- Corbet, P. S. 1980. Biology of Odonata. Annual Review of Ento- ent in an oligotrophic wetland ecosystem. Journal of the North mology 25:189–221. American Benthological Society 26:491–508. Corbet, P. S. 1999. Dragonflies: behavior and ecology of Odonata. Koch, K., C. Wagner, and G. Sahlén. 2014. Farmland versus Cornell University Press, Ithaca, New York. forest: comparing changes in Odonata species composition in Córdoba-Aguilar, A. (editor). 2008. Dragonflies and damselflies: western and eastern Sweden. Insect Conservation and Diver- model organisms for ecological and evolutionary research. sity 7:22–31. Oxford University Press, Oxford, UK. Kutcher, T. E., and J. T. Bried. 2014. Adult Odonata conservatism DeWan, A. A., and E. F. Zipkin. 2010. An integrated sampling as an indicator of freshwater wetland condition. Ecological In- and analysis approach for improved biodiversity monitoring. dicators 38:31–39. Environmental Management 45:1223–1230. Lemelin, R. H. 2007. Finding beauty in the dragon: the role of Dolný, A., F. Harabiš, and H. Mižičová. 2014. Home range, dragonflies in recreation and tourism. Journal of Ecotourism movement, and distribution patterns of the threatened drag- 6:139–145. onfly Sympetrum depressiusculum (Odonata: Libellulidae): a Le Viol, I., J. Mocq, R. Julliard, and C. Kerbiriou. 2009. The con- thousand times greater territory to protect? PLoS ONE 9: tribution of motorway stormwater retention ponds to the bio- e100408. diversity of aquatic macroinvertebrates. Biological Conser- Feindt, W., O. Fincke, and H. Hadrys. 2014. Still a one species vation 142:3163–3171. genus? Strong genetic diversification in the world’s largest Li, F., Y.-S. Kwon, M.-J. Bae, N. Chung, T.-S. Kwon, and Y.-S. living odonate, the Neotropical damselfly Megaloprepus cae- Park. 2014. Potential impacts of global warming on the di- rulatus. Conservation Genetics 15:469–481. versity and distribution of stream in South Korea. Garrison, R. W., and J. E. Hafernik. 1981. Population structure Conservation Biology 28:498–508. of the rare damselfly, Ischnura gemina (Kennedy) (Odonata: Loss, S. R., L. T. Terwilliger, and A. C. Peterson. 2011. Assisted ). Oecologia (Berlin) 48:377–384. colonization: integrating conservation techniques in the face Gee, J. H. R., B. D. Smith, K. M. Lee, and S. Wyn Griffiths. of climate change. Biological Conservation 142:92–100. 1997. The ecological basis of freshwater pond management Maxted, J. R., M. T. Barbour, J. Gerritsen, V. Poretti, N. Prim- for biodiversity. Aquatic Conservation: Marine and Freshwa- rose, A. Silva, D. Penrose, and R. Renfrow. 2000. Assessment ter Ecosystems 7:91–104. framework for mid-Atlantic coastal plain streams using ben- Hall, A. M., S. J. McCauley, and M.-J. Fortin. 2015. Recreational thic macroinvertebrates. Journal of the North American Ben- boating, landscape configuration, and local habitat structure thological Society 19:128–144. as drivers of odonate community composition in an island Mitsch, W. J., and J. G. Gosselink. 2007. Wetlands. 4th edition. setting. Insect Conservation and Diversity 8:31–42. John Wiley and Sons, New York. Hardersen, S., S. D. Wratten, and C. M. Frampton. 1999. Does Miura, T., R. M. Takahashi, and R. J. Stewart. 1990. Estimation carbaryl increase fluctuating asymmetry in damselflies under of absolute numbers of damselfly nymphs (Odonata: Coe- field conditions? A mesocosm experiment with Xanthocnemis nagrionidae) by dipper sampling in California rice fields with zealandica (Odonata: Zygoptera). Journal of Applied Ecology seasonal, spatial distributions and vegetation association. Jour- 36:534–543. nal of the American Mosquito Control Association 6:490–495. Harms, T. M., K. E. Kinkead, and S. J. Dinsmore. 2014. Evaluat- Monroe, E. M., and H. B. Britten. 2014. Conservation in Hine’s ing the effects of landscape configuration on site occupancy sight: the conservation genetics of the federally endangered and movement dynamics of odonates in Iowa. Journal of In- Hine’s emerald dragonfly, Somatochlora hineana. Journal of sect Conservation 18:307–315. Insect Conservation 18:353–363. Hart, L. A., M. B. Bowker, W. Tarboton, and C. T. Downs. 2014. Monteiro-Júnior, C. S., L. Juen, and N. Hamada. 2014. Effects of Species composition, distribution and habitat types of Odo- urbanization on stream habitats and associated adult dragon- nata in the iSimangaliso Wetland Park, KwaZulu-Natal, South fly and damselfly communities in central Brazilian Amazonia. Africa and the associated conservation implications. PLoS ONE 9: Landscape and Urban Planning 127:28–40. e92588. Moore, N. W. 1980. Lestes dryas Kirby—a declining species of Hassall, C., and D. J. Thompson. 2008. The impact of environ- dragonfly (Odonata) in need of conservation: notes on its mental warming on Odonata: a review. International Journal status and habitat in England and Ireland. Biological Con- of Odonatology 11:131–153. servation 17:143–148. Heiser, M., L. Dapporto, and T. Schmitt. 2014. Coupling impov- Morlando, S., S. J. Schmidt, and K. LoGiudice. 2012. Reduction erishment analysis and partitioning of beta diversity allows a in Lyme disease risk as an economic benefit of habitat resto- comprehensive description of Odonata biogeography in the ration. Restoration Ecology 20:498–504. Western Mediterranean. Organisms Diversity and Evolution New, T. R. (editor). 2012. Insect conservation: past, present and 14:203–214. prospects. Springer, Dordrecht, The Netherlands. Volume 34 September 2015 | 1031

Palmer, M. A. 1999. The application of biogeographical zona- lidae) with community participation in Yangon, Myanmar. Bul- tion and biodiversity assessment to the conservation of fresh- letin of Entomological Research 89:223–232. water habitats in Great Britain. Aquatic Conservation: Ma- Siegfried, B. D. 1993. Comparative toxicity of pyrethroid insec- rine and Freshwater Ecosystems 9:179–208. ticides to terrestrial and aquatic insects. Environmental Toxicol- Primack, R., H. Kobori, and S. Mori. 2000. Dragonfly pond ogy and Chemistry 12:1683–1689. restoration promotes conservation awareness in Japan. Con- Singh, R. K., R. C. Dhiman, and S. P. Singh. 2003. Laboratory servation Biology 14:1553–1554. studies on the predatory potential of dragonflynymphsonmos- Richter, B. D., D. P. Braun, M. A. Mendelson, and L. L. Master. quito larvae. Journal of Communicable Diseases 35:96–101. 1997. Threats to imperiled freshwater fauna. Conservation Stoks, R., S. Debecker, K. D. Van, and L. Janssens. 2015. Integrat- Biology 11:1081–1093. ing ecology and evolution in aquatic toxicology: insights from Rosset, V., J. P. Simaika, F. Arthaud, G. Bornette, M. J. Sam- damselflies. Freshwater Science 34:1032–1039. ways, B. Oertli, and D. Vallod. 2013. Comparative assessment Stoks, R., and A. Córdoba-Aguilar. 2012. Evolutionary ecology of scoring methods of the conservation value of biodiversity of Odonata: a complex life cycle perspective. Annual Review in ponds and small lakes. Aquatic Conservation: Marine and of Entomology 57:249–265. Freshwater Ecosystems 23:23–36. Strayer, D. L. 2006. Challenges for freshwater invertebrate con- Rosset, V., S. Angélibert, F. Arthaud, G. Bornette, J. Robin, A. servation. Journal of the North American Benthological So- Wezel, D. Vallod, and B. Oertli. 2014. Is eutrophication re- ciety 25:271–287. ally a major impairment for small waterbody biodiversity? Suhonen, J., E. Korkeamäki, J. Salmela, and M. Kuitunen. 2014. Journal of Applied Ecology 51:415–425. Risk of local extinction of Odonata freshwater habitat Samways, M. J. 1994. Insect conservation biology. Chapman generalists and specialists. Conservation Biology 28:783–789. and Hall, London, UK. White, E. L., P. D. Hunt, M. D. Schlesinger, J. D. Corser, and Samways, M. J. 2005. Insect diversity conservation. Cambridge P. G. deMaynadier. 2015. Prioritizing Odonata for conser- University Press, Cambridge, UK. vation action in the northeastern USA. Freshwater Science Samways, M. J. 2007. Insect conservation: a synthetic manage- 34:1079–1093. ment approach. Annual Review of Entomology 52:465–487. Yoshioka, A., Y. Miyazaki, Y. Sekizaki, S. Suda, T. Kadoya, and Sebastian, A., M. M. Sein, M. M. Thu, and P. S. Corbet. 1990. I. Washitani. 2014. A “lost biodiversity” approach to reveal- Suppression of Aedes aegypti (L.) (Diptera: Culicidae) using ing major anthropogenic threats to regional freshwater eco- augmentative release of dragonfly larvae (Odonata: Libellu- systems. Ecological Indicators 36:348–355.